Alice Giannetti, Paolo Monaco, Jess E. Caracuel, Jess M.

Soria &
Alfonso Ybenes
Functional morphology and ethology of decapod crustaceans
gathered by Thalassinoides branched burrows in
Mesozoic shallow water environments
Every part of a burrow has an important and specic
function with regard to the ecology and trophism of the
burrower. It goes from the guarantee of the oxygena-
tion and irrigation of the tunnels to the protection of the
organism or the storing of organic material for feeding
(Nickell & Atkinson, 1995). Regarding modern thalassi-
noidean burrows, presence of surface mounds, vertical
development of tunnels, geometry of the section of tun-
nels and horizontal galleries, presence of chambers and
of organic debris within the burrows, number and type
of apertures on the seaoor, presence of exhalant tunnels
and Y- or U-shaped burrow in vertical section, are gen-
erally considered as the most important characteristics
for ethological characterization of the burrow and of
the trace-maker (Nickell & Atkinson, 1995). Some of
these features can be recovered also in the trace fossil
Thalassinoides, usually considered a fossil counterpart
of modern thalassinoidean burrows, and they allow
one to formulate hypotheses about the trophism of the
trace maker. This can be particularly useful in those
cases in which body-fossil remains of the burrowers
are completely absent and a direct comparison between
the morphology of ancient and modern organisms is not
possible. Ecologic considerations are here applied to
Pliensbachian and Albian Thalassinoides developed in
the lagoonal deposits of the Trento carbonate platform
(Rotzo Member, Calcari Grigi Formation, northern
Italy) and in the outer shelf deposits of the Scaras For-
mation (Serra Gelada, Alicante Province southeastern
Spain), respectively. Different morphotypes of Tha-
lassinoides were identied in these two areas, mainly on
the basis of dimensions of the branches, diameter of the
turning chamber and development and geometry of the
maze. Even if Thalassinoides like burrows are nowadays
produced also by worms and shes, we consider crusta-
ceans to be the most probable trace makers of the studied
Thalassinoides. For a detailed discussion of the geologi-
cal setting, palaeonvironmental models, and ichnology
concerning the studied areas see Avanzini (1998), Gian-
netti (2004), Giannetti & Monaco (2002), Masetti et al.
(1998), Masetti & Posenato (2002), Monaco (2000a,
2000b), Monaco & Garassino (2001), Monaco & Gian-
netti (2001, 2002), Winterer & Bosellini (1981), and
Zempolich (1993) for the Calcari Grigi Formation, and
Caracuel et al. (2002), Castro (1998), Giannetti et al.
(2005), Monaco et al. (2005), Vilas et al. (1982), and
Ybenes (1996) for the Scaras Formation and refer-
ences therein.
In the Rotzo Member, three types of Thalassinoides
suevicus Rieth 1932 (Thalassinoides type I to type III,
from the smallest to the largest form) and another form
type of Thalassinoides isp. (type IV) were identied.
Thalassinoides suevicus type I is a small burrow, with
a diameter ranging from 2 cm to 5 cm. Branches are thin
and circular in section. Turning chambers are not par-
ticularly developed. Mazes are regular and geometrical
in shape and developed only on the horizontal plane.
Burrows are mud-lled and their surface is smooth. When
these burrows are in tiered with the other forms of Tha-
lassinoides, they occupy the upper tier.
In Thalassinoides suevicus type II, the diameter of
branches ranges from 5 cm to 10 cm and diameter of turn-
ing chambers ranges from 6 cm to 10 cm. Branches are
long and form geometrical, sometimes pentagonal mazes.
The external surface is smooth and lining is absent. Mud-
stone or bioclastic wackestone lls the burrows. Bioclasts
can be minutely fragmented and grouped in the internal
part of the burrow. This is the most frequently observed
type of Thalassinoides in the studied sections.
Thalassinoides suevicus type III is one of the largest
Thalassinoides form. Its diameter ranges between 10 cm
and 16 cm, and the diameter of the turning chamber is up
to 22 cm. Branches are squat with respect to the morphol-
ogy of the whole burrow and bifurcate at regular angles.
The outer surface is smooth and the lling is made of
bioclastic wackestones/packstones. Mottled concentra-
tions of minutely fragmented bioclasts can be present in
the central part of the burrow, in correspondence to the
lumen. This form always occupies the deepest tiers.
In Thalassinoides type IV the diameter of the branches
can reach 16 cm. The outer surface is mammillary, with
small mounds distributed without particular orientation
with respect to the burrow. Branches are short, subcircu-
lar in section and mazes are irregularly developed. On the
outer surface, crinoids and other bioclasts are frequently
grouped. Bioclastic wackestone lls the burrows.
Other cylindrical burrows, indicated as Thalassi-
noides? isp., exhibit simple tubes up to 80 cm in length,
are gently arched, 6 to 8 cm in diameter, and show short
aborted branches, without turning chambers. These
traces are very similar in horizontal plane view to those
illustrated by BROMLEY (1990). These atypical, branched
tunnels are very different from I-II-III-IV types described
above, and are analogous to some burrows of stomatopods
from the Seychelles Islands. For their morphological and
taphonomic characteristics these simple cylindrical tubes
49
3
RD
SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
are not closely related to burrows of shrimps or other
decapods, but resemble to those of carnivorous crusta-
ceans (stomatopods, Malacostraca), indicated as active
predators, now populating the shallow waters mainly of
tropical and subtropical seas.
In the Scaras Formation, we identied three types
of Thalassinoides suevicus Rieth 1932 (Thalassinoides
type A to type C) and another poorly branched form
(Thalassinoides type D), resembles the Thalassinoides ?
isp. described for the Rotzo Member, while the type IV is
very rare.
Thalassinoides suevicus type A is a small burrow,
with a maximum diameter of 5 cm. Turning chambers are
present but not particularly developed. It does not show
laterally developed mazes, but only burrows with short
branches developed only on the horizontal plane. Bur-
rows are mud-lled and their surface is smooth. Similarly
to the Rotzo examples, these burrows can be frequently
observed tiered with the other forms of Thalassinoides
and they are placed in the most supercial layers.
In Thalassinoides suevicus type B, branch diameter
ranges from 6 cm to 8 cm, turning chambers, can reach a
diameter of 9 cm. Regular mazes can be well developed
on the horizontal plane. The outer surface is smooth,
lining is absent and lling is homogenous and made up of
bioclastic wackestone.
Thalassinoides suevicus type C is the largest Thalassi-
noides form, with a mean diameter of around 8 cm, and
up to 12 cm in size for the turning chamber. When tier-
ing is present, this form is always located in the deepest
layers. Filling is homogeneous and made up of bioclastic
wackestone.
Thalassinoides isp. type D is a very large, horizontally
developed burrow, following an irregular zigzag path. It
is up to 1 m long, with a diameter ranging between 10
cm and 14 cm and a circular to slightly elliptical section.
It does not show true branching, but only short protru-
sions, extending in correspondence of the bending point
of the burrow. This trace resembles Thalassinoides? isp.
of Calcari Grigi. Filling is generally homogeneous and
is made up of ne-grained calcarenites or by bioclastic
wackestone/packstone, according to the lithology of the
host rock.
The following interpretation of the trophism of the
trace makers and of the ecological signicance of the
different Thalassinoides forms are based on the model
proposed by Nickell & Atkinson (1995; g. 3, p. 195)
for modern Thalassinoides-like burrows, considering the
main features common for both, modern and fossil bur-
rows. The 12 diagnostic features suggested by Nickell and
Atkinson (1995) and their ecological signicance are here
briey examined.
1) Surface mounds. They are produced by the trace-
maker removing sediment from the burrow and transport-
ing it to the surface. Surface mounds formed after the
construction of the burrow are indicative of deposit feed-
ing trophic mode. Cone-shaped, muddy mounds, locally
crossed by narrow vertical shafts are represented by a
few specimens in the Rotzo Member and are completely
absent in the Scaras Formation, maybe due to the higher
intensity of bottom currents.
2) Horizontal mazes (it replaces the tightly layered
lattice feature of Nickell & Atkinson, 1995). They sug-
gest the intense exploitation of the sediment by a deposit
feeder. Although all the studied Thalassinoides are hori-
zontally developed, this feature is particularly character-
istic of Thalassinoides type II, forming geometrical mazes
widely developed at the base of the beds. As suggested by
Suchanek et al. (1986), laterally extended mazes would
allow the trace makers to maximize the capture of organic
material when the substratum has a low nutritional value.
This probably could be related to different rates of nutri-
ent input within the sediment in the various phases of
evolution of the lagoon.
3) Deep burrows. Depth of the burrow development
within the substratum can indicate the importance of sur-
face sediments as a nutritional source for the trace-maker.
The deeper the maze is developed within the sediment, the
less the trace maker depends directly on organic materials
and current destruction present on the seaoor. Due to the
absence of vertical tunnels and to the vertical distribution
of the different Thalassinoides types within the beds, we
have hypothesized that the studied Thalassinoides, and in
particular the smallest forms, were probably developed
at a shallow depth within the substratum. This would
indicate a quite strong dependence of the trace makers on
organic material present on the seaoor (surface deposit
feeders or omnivorous scavengers). We must consider the
possibility that parts of burrows were eroded by bottom
currents and depth of burrowing within the substratum
could have been therefore underestimated. This consider-
ation is particularly important in the Serra Gelada section,
where burrows are developed in a relative high-energy
environment.
4) Sub-circular tunnel cross section. According to
Nickell & Atkinson (1995), sub-circular tunnels occur in
established burrows, after the phase of exploitation, and
are the result of crustacean movement. Oval or irregu-
lar shapes are less suitable for efcient water ow than
a perfectly circular cross-section (Nickell & Atkinson,
1995). The narrowing in cross section, as occurs in some
Thalassinoides? isp. (such as in some modern burrows of
stomatopods), is another characteristic which is not ideal
to water ow. According to this hypothesis, the occupant
of the burrow would be strongly dependent on the sur-
face sediments as a nutritional source and less to current
ows. With the exception of Thalassinoides type IV, all
the studied Thalassinoides both in the Calcari Grigi and
in the Scaras Formation have a sub-circular cross section
or narrowing, thus conrming the hypothesis suggested
by the other characters that the primary nutritional source
was the organic material present on the seaoor rather
than that transported passively by current ow within the
tunnels.
5) Chambered burrows. Chambers can have different
purposes: they can be used for sub-surface deposit feed-
ing or to store coarse grained material and fragmented
bioclasts, found within the substratum or coming from the
sea-oor. Another purpose maybe the change the direc-
tion of crustacean by 180, by performing a somersault
followed by a roll around the body axis, as observed in
modern examples of thalassinoideans. Therefore, the sub-
spherical shape of chambers maybe important to recognize
behaviour patterns, in the relationship with charateristics
of the substrate (Monaco & Giannetti, 2002). In Thalassi-
noides type III, type IV and secondarily in Thalassinoides
type II of the Calcari Grigi Formation and type B and C of
the Scaras Formation, turning chambers are particularly
50
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
developed and large. Stored coarse grained material such
as fragments of crinoids, coated grains and large bivalves,
algae and foraminifers were sometimes observed, in par-
ticular in Thalassinoides type IV.
6) Organic detritus in burrows. Organic detritus in bur-
rows can be the result of scavenging on the surface, but
also of passive falling of material within an open burrow.
In the fossil record, the different origin of organic detri-
tus can be sometimes identied through taphonomic and
sedimentologic criteria. In modern Thalassinoides-like
burrows, organic material is also used by the crustacean
to cultivate micro-organisms for nutritional purposes.
Organic detritus in burrows is particularly abundant in
Thalassinoides type III and type IV.
7) Oblique tunnels were very rarely recorded in the
studied sections (in some Thalassinoides ? isp.) and there-
fore they are not considered here.
8) Burrow openings, tubular tempestites. In the studied
sections, burrow openings can be directly observed only
rarely. Their existence can be deduced by the presence
of tubular tempestites inside the bed, but their number,
dimensions and density cannot be dened. The presence
of many burrow openings would indicate a continuous
and necessary contact of the trace maker with the seaoor,
for deposit feeding or scavenging.
9) Funnel-shaped openings. According to Nickell &
Atkinson (1995), the presence of funnel-shaped openings
would facilitate the capture of prey or any other nutri-
tional particles, as they fall down to the sides of the funnel
and into the burrow. In the studied section, this feature is
represented only by a specimen from the Rotzo Member.
The funnel is asymmetric in shape, the back being nearly
vertical and the front being attened probably due to the
continuous movement of the trace maker. The bottom of
the funnel is connected to a short vertical tunnel, which is
not directly related to a preserved Thalassinoides maze.
10) Narrow exhalant shaft. The presence of a narrow
exhalant shaft indicates the necessity of current genera-
tion. This feature was observed only in one cone-shaped
mound in the Rotzo Member, but its origin is uncertain.
11) U- or Y- shaped burrows in vertical section. This
feature, typical of many modern thalassinoidean burrow sys-
tems (Bromley, 1990), was not observed in the studied fossil
Thalassinoides and therefore it is not considered here.
12) Circular tunnel cross section. Their presence
would facilitate the water ow within and through the
burrow. Due to the importance of a constant water ow
through the burrow, circular tunnel cross sections are
present also in burrows produced by primarily deposit
feeders and therefore this cannot be considered diag-
nostic of a particular trophic mode (Nickell & Atkinson,
1995). In the studied sections this feature is only rarely
represented in Thalassinoides type II and in the largest
Thalassinoides type III, mainly where mazes are well
preserved and regularly developed.
In summary, features representing sediment processing
and storage of material for nutritional and maintenance
purposes are the most common in the Thalassinoides of
the Rotzo Member.
The only characteristic shared by the ve Thalassi-
noides forms is the horizontal development of the bur-
rows. Branches with a sub-circular section are typical
of Thalassinoides suevicus type I, type II, and type III.
Enlarged chambers are absent in Thalassinoides suevicus
type I and Thalassinoides? isp., present in type II and type
III and well developed in type IV.
According to the scheme proposed by Nickell &
Atkinson (1995), the most commonly observed features
in all the Thalassinoides of the Calcari Grigi Formation
indicate intense sediment processing. In Thalassinoides
suevicus type II, type III, and especially in the type IV,
storage of organic material (chambered burrows and
organic detritus in burrows, characters 5 and 6) is also
evidenced. The presence of organic detritus is represented
by coarse-grained to ne-grained crushed bioclasts within
Thalassinoides suevicus type II, type III, and type IV. It
indicates both biogenic sorting and storage of bioclas-
tic debris coming from the seaoor and fallen down or
trasported into the burrow. The local concentration of
coarse-grained fragmented shells in Thalassinoides type
IV, is probably due to the need of the trace maker to main-
tain other parts of burrow free from debris.
Vertical tunnels and apertures on the seaoor (char-
acters 8, 9 and 12) are very rarely observed and it is
sometimes impossible to state if vertical tunnels are not
preserved due to sedimentologic or diagenetic processes
or if they were really poorly developed in the burrows.
Actually, we have no evidences of deep burrows with a
U- or Y- shaped vertical morphology. We have hypoth-
esized, therefore, that mazes were burrowed on horizontal
planes only at shallow depth in the substratum. The few
evidences of vertical tunnels and of exhalant shafts are
difcult to relate to Thalassinoides mazes. The presence
of tubular tempestites inside the beds proves the existence
of direct apertures on the seaoor, even if only the hori-
zontal part of the lled burrows can be usually observed.
The only clearly observable vertical element has a circu-
lar section (character 12), but it is an isolated feature. This
tunnel closely resembles the burrows of modern crusta-
ceans and therefore we have considered it separately. In
the category of the vertical elements, we have considered
also inhalant and exhalant shafts, represented in the Rotzo
Member only by an unclear example. Actually, shafts
are so small that they could be produced also by worms.
These rare vertical elements indicate the presence of sus-
pension feeders, whose primary nutritional source is the
water column, and of surface deposit feeders. According
to the scheme proposed by Nickell & Atkinson (1995), all
the studied Thalassinoides belong to the category of the
sub-surface deposit feeders (searching for organic parti-
cles within the substratum) and secondarily to that of the
omnivorous scavengers (Vaugelas, 1990), which ingest
organic debris present on the seaoor and deriving from
other animals and algae.
Similar interpretations can be proposed for the Tha-
lassinoides studied in the Scaras Formation, even if
only four of the twelve characters described by Nickell
& Atkinson (1995), namely those related the horizontal
development of the burrow (characters 2-5), are well
preserved. The problem concerns the vertical develop-
ment of the burrows, because we have neither direct
nor indirect proofs of the existence of vertical tunnels
(except Ophiomorpha specimens of calcarenites at the
top of parasequences of Serra Gelada, see Monaco et
al., 2007) and of the type and number of apertures on
the seaoor. Anyway, we can hypothesize the pres-
ence at least of some vertical tunnels linking the big-
gest Thalassinoides, placed in the deepest tiers within
51
3
RD
SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
the sediment, to the seaoor. Intense bioturbation of
the more supercial tiers probably obliterated them.
All the studied Thalassinoides are horizontally devel-
oped (horizontal mazes, character 2) and show a sub-cir-
cular cross section (character 4), both characters indica-
tive of intense sediment processing.
Presence of enlarged chambers (as in Thalassinoides
suevicus type B and in Th. suevicus type C, character
5) could indicate storage of organic material for feed-
ing purposes or changes in the direction of locomotion.
Concentration of skeletal remains (character 6) is rarely
observed, due to the high bioclastic content of the host
rock and of the burrow lling. Because of the absence of
vertical tunnels, we cannot formulate hypotheses about
conditions of irrigation and importance of the water
column for nutritional purposes (characters 7-12). It is
evident that burrowers were closely dependent on nutri-
ents present within and on the substratum as the main
feeding source. Therefore, they can be included in the
trophic categories of the sub-surface deposit feeders and
secondarily in those of the surface deposit feeders and of
the omnivorous scavengers.
This is only a rst step in the ecological analysis of the
Thalassinoides trace fossils and of the trophism of their
trace-makers. However, an approach which considers the
functional morphology of the parts making up modern
Thalassinoides-like burrows can be useful for understand-
ing of the ecological meaning of ancient burrows produced
by extinct crustaceans and can give further information
for a more complete paleoenvironmental reconstruction
(Monaco et al., 2007). However, this method has to be
rened, elaborating a detailed model which compares dif-
ferent types of Thalassinoides in different environmental
contexts and considering also the functional morphology
of modern branched burrows produced not only by crus-
tacean decapods but also by other organisms.
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EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
Alice Giannetti - Geologisches Institut, Bonn Universitt, Nuallee 8, 53115 Bonn, Germany.
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3
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Memorie della Societ Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
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